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Metchnikovellids are highly specialized hyperparasites, which infect and reproduce inside gregarines (Apicomplexa) inhabiting marine invertebrates. Their phylogenetic affiliation was under constant discussion until recently, when analysis of the first near-complete metchnikovellid genome, that of Amphiamblys sp., placed it in a basal position with respect to most Microsporidia. Microsporidia are a highly diversified lineage of extremely reduced parasites related to Rozellida (Rozellosporidia = Rozellomycota = Cryptomycota) within the Holomycota clade of Opisthokonta. By sequencing DNA from a single-isolated infected gregarine cell we obtained an almost complete genome of a second metchnikovellid species, and the first one of a taxonomically described and well-documented species, Metchnikovella incurvata. Our phylogenomic analyses show that, despite being considerably divergent from each other, M. incurvata forms a monophyletic group with Amphiamplys sp., and confirm that metchnikovellids are one of the deep branches of Microsporidia. Comparative genomic analysis demonstrates that, like most Microsporidia, metchnikovellids lack mitochondrial genes involved in energy transduction and are thus incapable of synthesizing their own ATP via mitochondrial oxidative phosphorylation. They also lack the horizontally acquired ATP transporters widespread in most Microsporidia. We hypothesize that a family of mitochondrial carrier proteins evolved to transport ATP from the host into the metchnikovellid cell. We observe the progressive reduction of genes involved in DNA repair pathways along the evolutionary path of Microsporidia, which might explain, at least partly, the extremely high evolutionary rate of the most derived species. Our data also suggest that genome reduction and acquisition of novel genes co-occurred during the adaptation of Microsporidia to their hosts.

A selenid gregarine Ditrypanocystis sp. (Apicomplexa, Gregarinia, Selenidiidae), harboring the gut lumen of the oligochaete Enchytraeus albidus, was studied by light and electron microscopy. The trophozoite of Ditrypanocystis sp. is attached to the gut wall with its apical end to be anchored eventually between enterocytes in the crypts. Simultaneously, between the surfaces of the parasite and the host cell a peculiar contact is formed made of membranous channels and vesicles of unknown origin, the host cell surface in the contact area lacking cilia. The trophozoite becomes progressively enclosed within a parasitophorous vacuole made of layers of fused ciliar membranes of enterocytes. The fused cilia may be a source of membranes lining channels and vesicles of the contact area. Such a mode of parasitophorous arrangements has never been described before for gregarines, however, it bears a some likeness with that of the coccidian genus Cryptosporidium (similarity and differences being discussed). With regard to some molecular phylogeny constructions, claiming the "sister" relationship between gregarines and the coccidian genus Cryptosporidium (Carreno et al., 1999; Leander et al., 2003), this common feature in host-parasite relationships enabled us to put forward an idea of a possible evolutionary route from extracellularity of gregarines to intracellularity of coccidia, as exemplified by species of Cryptosporidium.

Several morphological forms (morphotypes) of Urospora chiridotae gamontes are found in White Sea holothuroid Chiridota laevis. All these morphotypes are differed by localization in the body of host, form and cytological features. The gregarines are situated in several host biotopes, such as blood vessels, intestine and mesenteries. In the blood vessels elongate skittle-like cells supplied with long thin cytopillia are observed. On the external surface of the intestine spherical gregarines are found. These parasites commonly covered with one layer of coelomic epithelium's cells. In some holothuria intratissue spherical cells of parasites located in intestinal epithelium are presented. Both of these types of parasites lack cytopillia, and folds or ridges on its surface. On different mesenteries, connections between intestine and body wall, and also on intestine elongate ounce-shaped cells and gamontocysts are observed. These cells are situated on the apices of finger-like processes of the intestine and mesenteries surface. Ounce-shaped gregarines have cytopillia shorter than in skittle-like gregarines. The differences between morphotypes of Urospora chiridotae are probably caused by different environmental conditions. In the narrow rift of blood vessel elongate cells are developed. The cytopillia may serve for making more or less wide space around gregarines, which is necessary for food uptake. Spherical cells surrounded by host's cells and have the form typical for tissue parasites. In the wide coelomic cavity where convection of liquid proceeds better than in blood vessel, ounce-shaped gregarines with short cytopillia are developed. We found only typical for Urospora chiridotae ovoid oocysts with dissimilar ends, anterior collar and spine-like posterior end. Thus, the all above-mentioned morphotypes undoubtedly belong to the same species. The relationships between defense host cells and the different morphotypes of trophozoites are variable.